Ann,cool. Fennici17:107-112. 1980.
The status, structure and behaviour of populations of the wolf (Canis I. lupus L.) along the Fenno-Soviet border1
Erkki Pulliainen
Pulliainen, E. 1980: The status, structure and behaviour of populations of the wolf (Canis I. lupus L.) along the Fenno-Soviet border. - Ann. Zool. Fennici 17:107-112.
Since 1950 Finland has received two expansions of the wolf ( Canis l. lupus) from the east, mainly from Soviet Karelia into Finnish Northern Karelia and Kainuu, the first in 1959-63 and the second beginning in 1975. Before and during both expansions most of the wolves wandering in Finland were males, but as the breeding population approached the frontier the excess of males decreased. Groups reaching western and southern Finland (mainly along certain specific migration routes) usually consisted of one to four wolves, but in eastern Finland some packs were larger, and in 1969-79 pups aged 4-10 months were also seen. From studies carried out in Soviet Karelia and records of wolves crossing the Fenno-Soviet border it appears that under present-day conditions the saturation point of the Soviet Karelian wolf population is roughly five to seven individuals per 2 1000 km • Continuous hunting of a wolf population disturbs its self-regulation mechanism and enables it to attain its potential maximum productivity, which explains the rapid growth of populations in Soviet Karelia and the Leningrad Region of the USSR in the 1970s. In January 1979 there were about 100 wolves in Finland.
Erkki Pulliainen, Department of <::,oology, University of Oulu, Kasarmintie 8, SF-90100 Oulu JO, Finland.
I. Introduction The purpose of the present paper is to provide records on the status, structure and behaviour of At the beginning of the 19th century the range wolf populations along the Fenno-Soviet border. of the wolf (Canis l. lupus L.) included the whole of Finland. The decline began on the Aland Islands, where the last wolf was killed in 1844 2. Material and methods (Salovaara 1930), but in the 1880s wolves were still to be found in all parts of the Finnish When studying the wolf expansion of 1959-63, I was mainland. Thereafter, the decli.ne was rapid, and greatly assisted by the Finnish Border Patrol Establish by 1900 the species bred only in the eastern and ment (Pulliainen 1965), and these experiences led me to northern parts of the country (Palmen 1913). suggest that the daily patrols along the frontiers between Finland and its neighbouring countries could record In the Scandinavian Peninsula the wolf popu every crossing of the frontier by the large predators and lation had been decimated some decades earlier estimate or calculate the numbers of these mammals in than in Finland, although the remnants were the areas under their surveillance three times a year. This able to survive in northern Scandinavia for a proposal was accepted by the commander of the Border considerable time with a surprisingly small Patrol Establishment, and data from this observation line population base (Haglund 1973). As far as 2574 km long are now available for a period of over 11 years (1968-1979). The records made during this time Finland is concerned this base has been the wolf show a total of 4195 crossings of the frontier by populations inhabiting the Kola Peninsula, Soviet wolves. Karelia and the surroundings of Lake Ladoga. Wolf tracks are easy to observe and identify in snow. In the northernmost parts of Finland the snow disappears in late May or early June and may appear by the end of 1 Report No. 100 from the Varrio Subarctic Research September, or more normally in October, while in the Station of the University of Helsinki. south-east it may last only 3 or 4 months. This difference 108 Erkki Pulliainen
must be kept in mind when assessing the crossing data, 3. Results although even in snowless conditions wolf tracks can be identified in sandy, wet or muddy ground, for instance_. It 3.1. Finnish wolf populations should be noted that the members of the Border Patrol Establishment are taught during their preliminary train Of the 4023 crossings of the frontier by wolves ing to recognize the tracks of wolves and other large recorded in 1968-78 by the Border Patrol predators. Establishment 4007 (99.6 %) took place across the The Russians follow the trends in the populations of the border between Finland and the USSR, 14 (0.3 large predators by skiing along observation lines in %) on the Norwegian frontier and two (0.1 %) on different parts of the country and counting the numbers of the Swedish frontier. During 1968-72 the num tracks of these animals crossing ·the lines (Danilov et al. ber of crossings varied between 70 and 115, in 1978). The results are then expressed as the numbers of tracks per 10 km of observation line. 1973 the records showed 192 crossings, and there was then a very sharp rise, from 129 in 197 4 to It is interesting to compare the Finnish data with the 1259 in 1977 (Fig. 2). This rise took place corresponding Soviet Karelian data. Fig. I shows the total numbers of crossings of the frontier by wolves especially in Northern Karelia (Lieksa and recorded in Finnish Northern Karelia and the numbers of Ilomantsi) and southern Kainuu (Kuhmo), alth tracks/IO km found in the corresponding Soviet Karelian ough in 1976 and 1977 wolves were recorded all area in 1968-76. Upward trends are clearly visible in along the border between Finland and the USSR both sets of data, and can be explained in ten of a';i -in (Fig. 3). At that time no rise was shown by the increase in the numbers of wolves in Soviet Karelia from figures for the Norwegian or Swedish frontiers, 1971 onwards, leading to an expansion into Finnish territory after 197 4. although a few wolves did enter Swedish Lap land from Finnish Lapland. The crossing data The crossing data give a reliable picture of the relative and other observations made indicate that 1977 changes in the mobility of wolves (see also Pulliainen 1979b), but do not reveal the actual number of animals may represent the peak year for the present involved, for one individual may cross the frontier several expansion of wolves from the east, for the total times during a year. Nor is it reasonable to calculate rates number of crossings was smaller in 1978, and this of immigration or emigration from these figures, as few downward trend has continued during the first 5 observations can be made during the snowless season, and months of 1979. possibly the wolves return to the USSR to breed at that The patrols have estimated the numbers of time of year. wolves three times a year (1 January, 1 June and Information on all wolves killed and found dead in 1 October) over the 11-year period within a zone Finland was sought through every available channel of enquiry.
1200
eoo 1000
800 ..: 40 ► ., ., " 600 'll: u '0 C 0-4" ..°' 20 ll ~--1· 400 0>'---~-- _ _.__·----- _ _.__ _ _._- _ _.__~-~-~ 0 11169 1971 1973 1975 200 Fig. 1. The total numbers of crossings of the frontier in Finnish Northern Karelia (left scale; solid line) and the mean numbers of crossings of observation lines by wolves 1978 in the adjacent Soviet Karelia (right scale; dotted line; 1968 1970 1972 1974 1976 Danilov et al. 1978). A star indicates the saturation point Fig. 2. The total numbers of crossings of the Finnish of the wolf population of this part of the Karelian ASSR frontier by wolves recorded by the Border Patrol Estab (see also text). lishment in 1968-78 (see also text). Ann. Zool. Fennici 17. 1980 109
The sexing of 123 dead wolves in Finland in 1969-1979 showed that 65.8 % were males, a disparity which is statistically highly significant. The 48 wolves examined in 1968-75 included 38 males (almost 80 %), and correspondingly the 75 wolves examined in 1976-79 included 43 males (57.3 %), there being still a slight, although statistically non-significant, excess of males. Table l shows the sex ratios of wolves killed in different parts of Finland in 1948-63 and 1969-79. The wolf is known to have bred in the north of Finland (Lapland) in 1948-53, and at that time the 25 dead individuals included nine females. In 1969-79 almost all the wolves moving in this area were killed by reindeer owners, and among nine individuals sexed there were seven males. Both the district between Savukoski and Kuh mo and that of Northern Karelia lie along the border between Finland and the USSR (Fig. 5). The expansion of the wolf from the east, which reached Northern Karelia and the southern part of the Kuhmo-Savukoski district in 1958-63, led to an even sex ratio in the former area but an excess of males in the latter. In 1969-79 the Savukoski-Kuhmo district also had a statistically Fig. 3. The numbers (calculated per 100 km of frontier) of crossings of the different parts of the Finnish frontier by significant excess of males. In Northern Karelia wolves recorded by the Border Patrol Establishment in there were five males and one female among the 1968-78. dead wolves encountered in 1972-75, but an even ratio of 16: 15 in 1976-79. Excesses of males among the dead wolves were found in 1948-63 at least 20 km wide adjacent to the frontier. The and l 969-79 in the other parts of Finland where most reliable estimates are naturally those based wandering wolves occurred. on observations in the snowy season, i.e. l Among the 29 wolf pups aged 4-10 months January. These show that the majority of wolves killed in Finland in 1969-79 (Table 2), there occurred in the vicinity of the frontier between Finland and the USSR (Fig. 4), the numbers varying between 6 and 24 in 1969-75, but □ USSR increasing from 1976 to 1978. The total figure 80 reported for l January 1978 was somewhere isl NORWAY between 77 and 89, but on l January 1979 it was SWEDEN again smaller (between 63 and 70). The majority of the "Finnish" wolves taken 60 • into account in these estimations have been moving in the vicinity of the border between Finland and the USSR in units of l to about IO 40 individuals, and may have crossed the border almost daily. When pursued on the Finnish side, they retreated to the Russian side. In the inner parts of Finland wolves have also been seen moving in units of 1-4. They use the old migration routes (Fig. 5 ), and since 1970 in dividuals have been seen in western and southern 1969 -71 -73 -75 -77 -79 Finland. The total information available in Fig. 4. The numbers of wolves in the vicinity of the dicates some 30 wandering wolves in January Finnish frontier on I January 1969-79 according to the 1979. Thus at the beginning of 1979 Finland had estimate of the Border Patrol Establishment (see also about 100 wolves. text). 110 Erkki Pulliainen
Table I. Sex ratios of wolves killed in Finland in 1948-63 and 1969-79. Area 1948-63 1969-79 c:!% n ,:!•70· n
Lapland 64.0 25 77.8 9 The Savukoski-Kuhmo district 72.3 47 67.2 58 Northern Karelia 46.9 49 56.8 37 The other parts of Finland 88.9 18 73.6 19
the late 1940s intensive clear-cutting was com menced in these forests, and the conifers were replaced by deciduous trees, which offered food for the moose populations and enabled these to increase greatly. At the same time reindeer hus bandry was discontinued in the area, and the semi-domestic reindeer (R . t. tarandus L.) re turned to the wild state, while the wild forest reindeer (R. t. fennicus Lonnberg) were no longer hunted. Thus there was abundant food for the wolves, which could move freely along the forest roads, trails of ungulates, and other paths. Consequently, the wolf population expanded northwards at the same time both in Karelia and Fig. 5. The main migration routes of wolves in Finland, in the adjacent Arkangelsk area (see also numbers of crossings of observation lines by wolves in the Semenov 1976). Karelian ASSR in winter (according to Danilov et al. Danilov et al. (1978) drew attention to the 1978; abbreviations: I = 0.19 or less, 2 = 0.20-0.29, threefold increase in the Karelian wolf popu 3 = 0.30-0.39, 4 = 0.40-0.49 and 5 = 0.50 or more tracks/ IO km of observation line) and the Finnish study lation from 1966-69 (0.2 tracks/ IO km) to areas mentioned in Table I (A = Lapland, B = the Savu 1973-76 (0. 7 tracks/ IO km), attributing this to koski-Kuhmo district, C = Northern Karelia and D = the two factors, the improved food situation men other parts of Finland). tioned above and the reduced control during the years when small numbers of wolves were recorded. · was a slight, but statistically non-significant, Bibikov (1973) reported that there were 300 excess of males (58.6 %). There are probably only wolves in Karelia in the early 1970s. These two sets of data, taken together, can be used to two cases (Nos. XI and XVI) in which the figures estimate that at the time of the maximum in 1976 represent a complete litter. there may have been 800-900 wolves in Soviet Karelia. During the 1978 hunting season 151 3.2. Soviet Union wolf populations wolves were killed in the area (official statistics). Bibikov (1973) further stated that at that time According to Danilov et al. (1978), the mean there were 140 wolves in the Leningrad area, number of wolf tracks per IO km in the whole of where the population probably also increased, Soviet Karelia in 1961-76 was 0.4. The popula for, according to the local authorities, there were tion was largest (0.5-0.6 tracks/ IO km) in the some 500 in that area in 1978. The official goal is southern and central parts of this area (Fig. 5 ), to reduce the number to 150 in a forest area of 2 which are characterized by abundant popu about 60 000 km • lations of moose (Alces alces L.) and reindeer In 1973 Bibikov reported that there were 30 ( Rangifer tarandus L.); in the northeastern part wolves in the Kola Peninsula, inhabiting its there are fewer moose and reindeer, and also western and southern parts. No later reports are fewer wolves (0.2-0.3 tracks/ IO km). available, but observations made in eastern Danilov et al. (1978) report that before 1940 Finnish Lapland can tell us something about the wolves were rare in northern Soviet Karelia (see later history of this population. also Marvin 1959), the few individuals inhabiting Thus there are breeding wolf populations cultivated areas and the coasts of the White Sea. beyond Finland's eastern frontier throughout its Practically no wolves occurred in the taiga. In le~gth, although the western edges of these Ann. Zoo!. Fennici 17. 1980 111
Table 2. Sex ratios of wild wolf pups in Finland according to kill statistics.
Date of Litter Locality where a wolf died Pups Other members of death no. 6 ~ the pack killed
IX.1969 1• Suomussalmi, Raate 0 Xl.1969- 11 .1970 11• Kuhmo, Lentua-Kalliojoki I I III.1972 III Lieksa, Lehmivaara I 0 Xl.1972 IV Savukoski, Lipakkaselka 0 I Xl.1972 V Salla, Onkamojarvi 2 0 1.1973 VI Kitee, Papinnien1i I 0 11.1973 VII llomantsi 0 I X.1973 VIII Salla, Kulvakkovaara 2 0 adult 6 XI.1974 IX Kuusamo, Liikanen I 0 11.1976 X Nurmes I 0 X-XII.1976 XI Salla, Niemela-Selkala I 2 adult 17 12 •may belong to the same litter. populations may or may not reach Finnish communication), but if the a-male is removed, territory (Fig. 3). all the mature females give birth to pups (Ginsburg, pers. communication). The a-pair is chiefly responsible for the care of the young, and 4. Discussion it is therefore most vulnerable (of the adults) to the In northern Europe the wolf has for decades hunter (see also Table 2). Wolfhuntingthusallows occupied a greatly restricted range. Within the the potential maximum productivity of the pack area of western culture the species has been to be realized (see also Mech 1970). The wolf exterminated almost entirely, while in eastern populations of Soviet Karelia and the Leningrad Europe its populations have made rapid growth. region have been hunted continuously, although The eastern frontier of Finland forms a border at varying intensities (Danilov et al. 1978). between these two cultures. This dichotomy is A wolf population at saturation point natur reflected very strikingly in the occurrence of the ally disperses in directions where there are no species on the frontiers of Finland during the last barriers and into suitable empty territories. In decade; very few crossing have been recorded on the case of Soviet Karelia and the Leningrad the Swedish and Norwegian frontiers, while region the latter were available in Finland, more than 4000 observations have been made on which is part of the former range of the species. the eastern frontier. The crossing data also The wolf was protected in the southern half of appeared to agree fairly well with our knowledge Finland from September 1973 until the end of of the sizes of the wolf populations beyond the 1977, and events showed that the species would borders of Finland. probably very soon regain its former territories, In Soviet Karelia and the Leningrad region despite the drastic changes made by man in the the· increase in the wolf populations has been structure of the landscape. Indeed, wolves might rapid, even threefold during less than a decade. successfully inhabit even the most settled areas in In areas where there is no human impact on the southern Finland, if they were not faced with the wolf population, e.g. on Isle Royale, such in hazards of our "civilized world" in the form of creases do not seem to occur (see Mech 1970, busy roads and unintentional disturbance by Peterson 1977, Allen 1979). The reason for this is man (see Pulliainen 1979a). probably the self-regulation mechanism of the Bibikov (1973) estimates that the 300 wolves in wolf population, for the suppression exerted by Soviet Karelia represent a density of 2.5 wolves 2 the a-pair is so great that the other mature per 1000 km • Since expansion can be regarded females of the pack do not produce offspring. as an expression of population pressure in a This has been verified in nature (Haber 1977) population at saturation point, recent observa and in captivity (Zimen 1976, G_insburg, pers. tions on the increase in the Soviet Karelian wolf 112 Erkki Pulliainen population and the commencement of a powerful killed, and expansion was thus blocked (Pulliai expansion into Finnish Northern Karelia allow nen 1965). Before and during that expansion us to estimate (see Figs. 1 and 3) that the most of the wandering wolves were males, but as saturation point of the wolf population under the the breeding population approached the frontier conditions prevailing at present in Soviet Karelia the excess of males decreased. The same trend in is roughly 5-7 wolves per 1000 km2. In the sex ratios has been recorded during the present European part of the USSR the wolf populations expansion from Soviet Karelia into Finnish have reached even higher densities (IO or more Northern Karelia (Table 1), and a similar 2 ind./1000 km ) in the more southerly regions blocking of the expansion is now in progress, for (Bibikov 1973). . at least 72 wolves have been killed in _Finland in Concerning the behaviour of a wolf population 1977-79. at saturation point under present-day conditions In 1959-79 wolf packs comprising more than in Soviet Karelia, one might speculate that the five individuals have been observed only in the use of forest roads and the trails of ungulates as easternmost part of Finland (Pulliainen 1965, pathways in deep, soft snow (see also Danilov et present data). Units which have wandered to the al. 1978) would lead to increased numbers of western and southern parts of the country have encounters between wolf units (lone wolves and consisted of one to four individuals. Most of these packs) and thus to increased mobility, which wandering groups were probably formed by low may be manifested both in an increased number ranking adults and juveniles from the packs in of crossings of the observation line (e.g. the the breeding area, which show a greater tend border between Finland and the USSR), and ency to emigrate than do the other members of also in a strong tendency for the population to the packs (see also Zimen 1976). It is worth disperse. I have taken these aspects into account noting that no return of such wandering indi in estimating the size at which the Soviet viduals has been recorded so far, which suggest Karelian wolf population reached saturation that (a) this is real dispersal of a population(s) point. from a saturated area and (b) the wandering The majority of the wolves which crossed into individuals have found suitable habitats and Finland from Soviet Karelia in 1959-63 were enough available food in their new areas. References Allen, D. L. 1979: Wolves of Minong. Their vital role Mech, L. D. 1970: The wolf: the ecology and behavior in a wild community. - 499 pp. Boston. of an endangered species. - 384 pp. Garden City, Bibikov, D. I. 1973: The wolf in the USSR. - Paper New York. presented at the Technical meeting on the wolf in Palmen, J. A. 1913: Bar, Wolf und Luchs in Finnland. - Europe, IUCN/ WWF, Stockholm, September Zoo!. Beobachter 54 (3): 1-6. 5-6 1973, 9 pp. Peterson, R. 0 . 1977: Wolf ecology and prey relationships Danilov, P. I., lvanter, E. V., Belkin, V. V. & Niko in Isle Royale. - National Park Service Scientifics laevskij, A. A. ()];aHHJioB, II. IL, liBaHTep, Monograph Series 11 : l-210. E . B ., Bemnrn, B . B. & H1rnoJiaeac1rntt, Pulliainen, E. 1965: Studies on the wolf (Canis lupus L.) A. A.) 1978: HaMeHeHH&a 'IHCJieHHOCTH 0XOT in Finland. - Ann. Zoo!. Fennici 2:215-259. HH'IHX auepett KapeJIHH no MaTepHaJiaM 3HM -»- 1979 a: Ecology of the wolf in the settled areas HHX MaprupyTHblX y'leTOB. - In: lvanter, E. V. of Finland. - In: Klinghammer, E. (ed.): The (ed.):